Bulletin of the California Lichen Society 2009 16-1

BULLETIN OF THE CALIFORNIA LICHEN SOCIETY 16 1, 2009 McGee – Spore-PrintsI photograph and measure the spores in water first, and later may mount the spores in other media for other treat

Bulletin

of the

California Lichen Society

Volume 16 No 1 Summer 2009

The California Lichen Society seeks to promote the appreciation, conservation and study oflichens The interests of the Society include the entire western part of the continent, although thefocus is on California Dues categories (in $US per year): Student and fixed income - $10,Regular - $20 ($25 for foreign members), Family - $25, Sponsor and Libraries - $35, Donor -

$50, Benefactor - $100 and Life Membership - $500 (one time) payable to the California LichenSociety, PO Box 7775 #21135 , San Francisco, California 94120-7775 Members receive theBulletin and notices of meetings, field trips, lectures and workshops

Board Members of the California Lichen Society:

President: Erin Martin, shastalichens gmail.com

Vice President: Michelle Caisse

Secretary: Patti Patterson

Treasurer: Cheryl Beyer

Committees of the California Lichen Society:

Education/Outreach: Erin Martin, chairperson

Poster/Mini Guides: Janet Doell, chairperson

Events/field trips/workshops: Judy Robertson, chairperson

The Bulletin of the California Lichen Society (ISSN 1093-9148) is edited by Tom Carlberg,tcarlberg7 yahoo.com The Bulletin has a review committee including Larry St Clair, ShirleyTucker, William Sanders, and Richard Moe, and is produced by Eric Peterson The Bulletinwelcomes manuscripts on technical topics in lichenology relating to western North America and

on conservation of the lichens, as well as news of lichenologists and their activities The best way

to submit manuscripts is by e-mail attachments or on a CD in the format of a major wordprocessor (DOC or RTF preferred) Submit a file without paragraph formatting; do include italics

or underlining for scientific names Figures may be submitted electronically or in hard copy.Figures submitted electronically should provide a resolution of 300 pixels-per-inch (600minimum for line drawings in JPEG format); hard copy figures may be submitted as linedrawings, unmounted black and white glossy photos or 35mm negatives or slides (B&W orcolor) Email submissions of figures are limited to 10 MB per email, but large files may be splitacross several emails or other arrangements can be made Contact the Production Editor, EricPeterson, at eric theothersideofthenet.com for details of submitting illustrations or other largefiles A review process is followed Nomenclature follows Esslinger cumulative checklist on-line

at http://www.ndsu.nodak.edu/instruct/esslinge/chcklst/chcklst7.htm The editors may substituteabbreviations of author’s names, as appropriate, from R.K Brummitt and C.E Powell, Authors ofPlant Names, Royal Botanic Gardens, Kew, 1992 Instructions to authors will soon be available

on the Society’s web site (below) Style follows this issue Electronic reprints in PDF format will

be emailed to the lead author at no cost

The deadline for submitting material for the Winter 2009 CALS Bulletin is October 31 2009.

The California Lichen Society is online at http://CaliforniaLichens.org and has email discussionsthrough http://groups.yahoo.com/group/CaliforniaLichens

Volume 16 (1) of the Bulletin was issued 31 August 2009

Front cover: Pannaria rubiginosa and captured spores (see paper on page 1) Photography by

Mikki McGee

Bulletin of the California Lichen Society

VOLUME 16 NO 1 SUMMER 2009

Spore-Printing Lichens

Mikki McGee

8 Visitacion Ave #10Brisbane, CA 94005 mikkimc earthlink.netSexual spores of lichens are important to the

lichen-fungus as well as to the person identifying

lichens, or classifying them The person identifying

or classifying lichens wishes to have clear

unobstructed or unobscured view of as many spores

as possible Their sizes and shapes and septation are

used in distinguishing one species from the other

For the fungus, spores are arguably less important as

propagules for starting new thalli than as a sexual

means for maintaining sexual or genetic diversity in

the thallus and the population

Finding enough spores that are clearly visible to

identify the lichen can be difficult Tissues may be

overlaying or underlying the free spores found,

making them less clearly visible And spores still in

asci are possibly distorted by the pressures of the

ascus

Mycologists studying mushroom fungi make a

regular practice of trying to get a collection of spores

on a piece of paper by laying the mushroom on paper

in a moist environment (covered by a dish) for an

hour or so The spores falling onto the paper are then

examined for dry color, and examined

microscopically for size and form The paper on

which the print is made often stays with the specimen

for later examination Ascus-bearing larger fungi

(dime to dinner plate size) sometimes are treated the

same as the more common basidiocarps

The fresh smaller ascocarps of lichens may also

produce deposits of spores which can serve some of

the same processes, when collected on cover glasses

I use 12x12mm cover glasses routinely, and often

easily collect more than enough spores to

characterize the spores and the lichen, and they are

free of any tissues that might obscure them

The method is simple, but does not work for

most of the material already long dead in herbaria It

may be used successfully on material that has notbeen forcibly air dried or frozen Lichens fresh fromthe field are best If there are several specimens, one

is selected If only one is available, then it may besubdivided to produce one fragment for sporeprinting, or the entire collection used, at thediscretion of the worker A single fruiting body may

be spore printed successfully, if it is fresh and aliveand undepleted on collection

The lichen is placed in a petri dish, and soakedwith distilled water (Tap water is generally treatedwith chemicals that may harm the lichen, and bottledwater often contains many more bacteria andprotozoa than tap water.) When the lichen is wellsoaked, excess water is poured off, and a cover slip islaid over several fruiting bodies If the specimen isquite small, a folded wad of wet paper towel may beenclosed in the dish to maintain moisture (Figure 1)

It is common for water to seal the cover glass to thelichen - it holds the cover glass in place

The asci, on discharge, forcibly shoot the sporessome distance, and the spores being sticky attach tothe cover glass After an hour or so, I examine thecover glass in water on a slide, to look for spores Inthe meantime, I may be preparing other parts of thesame collection, or having coffee

Water is the first choice of mounting media Thespores so examined may still show their halos well, ifthey have them The halo, being of weak gelatin,once dried and shrunk may not re-expand later Ifone wishes to preserve the halo for later examination,transferring the cover-glass after measurement to apreserving mountant such as Hantsch's fluid mayserve I also use formalin-alcohol-acetic acid (FAA)

to kill, fix and preserve spores and other things forless distorted later viewing than with other methods.But the first measurement is always in water

BULLETIN OF THE CALIFORNIA LICHEN SOCIETY 16 (1), 2009 McGee – Spore-Prints

I photograph and measure the spores in water

first, and later may mount the spores in other media

for other treatments If several cover glasses are used,

one can treat each one differently - one for staining in

iodine, or one to Hantsch's, eliminating oil droplets to

see the septa more clearly The treated spores are then

photographed and measured again, to see and record

the effect of the treatments Some of the spores may

detach from the cover slip and be found on the slide

when the cover slip is lifted, making two

preservations possible, as well

Some spores, as of Teloschistes, Xanthoria and

Caloplaca, are routinely heat treated to kill and dry

them, for comparison to herbarium type material

This is easily done on the cover slip after the initial

examination For my work, I want to see them fresh,

as well It is interesting to see the fresh viable spores,

as well as seeing the mummies the taxonomists love

Hantsch's Fluid:

1 part(ml) glycerin with 2 parts water are mixed

as stock solution One drop of 95% alcohol is added

to one drop of this stock for use, yielding two drops

of 1,2,3 glycerin-water-alcohol, which on evaporation

of the alcohol and water can make permanent ringedslides of one 24mm or two 12mm cover glasses.Preserved slides are placed in packets after ringing

Formalin-Alcohol-Acetic Acid (FAA):

This is a standard preservative, which for fungiand lichens is usually mixed 10 parts 40% formalin,

50 parts 95% Alcohol, 5 parts glacial acetic acid;with 35 parts of water (Total 100 parts.)

The glacial acetic acid can be adjusted too: from

2 parts to 6 parts, (2%-6% final acidity) to reduceshrinkage, or swelling from formalin and alcohol.Adjust by exchanging water for acid, or vice verse.(Email me for answers to questions.)

Figure 1 Pannaria rubiginosa (from the San Bruno

Mountain field trip, reported elsewhere in this issue)

prepared for spore printing Note the paper toweling

beneath the lichen The edges of the cover slip are just

visible Printed in color on front cover

Figure 2 Spores captured using the methods inthis article Printed in color on front cover

BULLETIN OF THE CALIFORNIA LICHEN SOCIETY 16 (1), 2009 Tucker – New Reports

New Reports and Comments on California Lichens

Shirley TuckerSanta Barbara Botanic GardenSanta Barbara, CAtucker lifesci.ucsb.eduNew additions to the California lichen flora are

published continually, and some of these merit

comments to stimulate searching and collecting in the

state Most are not in the most recent published

checklist for the United States (Esslinger & Egan

1995), but all are in the online checklist by Esslinger

(2008) I add these to my manuscript of the

California catalogue (Tucker & Ryan, 2006) and

eventually these additions will be inserted in the

online version of the catalogue Meanwhile, the

following deserve attention from collectors

Byssoloma leucoblepharum (Nyl.) Vainio

I found this crust once on bark in Mendocino

County (S Tucker 35280, Pygmy forest, Mitchell

Creek Drive near Fort Bragg, SBBG) The apothecia

are unmistakable because of the “byssoid” margin,

like a fluffy halo of outward-extending colorless

hyphae around the dark disk There is a photograph

of this crust in Brodo et al 2001 (p.192) showing the

purplish-brown disk and pale margin, although the

magnification is not great enough to show the fluffy

margin It is labeled as B meadii, but that species has

pale yellow disks and is in southeastern U S Another

species, B marginatum, has been found in Sequoia

sempervirens treetops of Humboldt County by

Williams & Sillett (2007) The minute apothecia of

this species are pale greenish-gray and lack a byssoid

margin

An unusual lichen niche on the Pacific

Northwest coast was mentioned by Williams & Sillett

(2007) and more recently emphasized by Spribille et

al (2009) Unusual minute lichen crusts occur on

stems of ericaceous evergreen shrubs such as species

of Vaccinium and Rhododendron, close to ground

level in rain forests Collectors in northern CA could

add new reports to the California lichen flora by

searching this niche

Ptychographa xylographoides Nyl

McCune (1997) described this taxon from

Oregon; Williams & Sillett (2007) climbed to the

tops of redwoods in Humboldt County and found an

amazing flora of lichens, mosses and angiosperms

growing there They found 183 lichen species, 50

bryophytes, and 49 species of vascular plants Thearticle is well worth reading, as it not only describesthe rope-climbing techniques used to access thetreetops, but also discusses in detail the many andvaried niches where lichens may grow – on live anddead foliage, large vs small branches and twigs, barewood, soil and plant debris lodged in branch

crotches, and tree litter Ptychographa is found on

dead wood, especially in conifer forests, the same

habitat as the similar Xylographa McCune (1997)

mentioned a means of field identification:

Ptychographa resembles a Graphis or Opegrapha in

having black lirellae (elongate apothecia) but has amiddle ridge separating two parallel elongatehymenia The spores are colorless unicells, like those

of Xylographa but quite unlike the septate spores of either Graphis or Opegrapha

Other lichens new to California found byWilliams & Sillett (2007) include a new species of

pin-lichen, Calicium sequoiae (Williams & Tibell 2008), three newly reported species of Micarea

(determined by Brian Coppins), and the first state

report of Segestria leptalea, a pyrenolichen with tiny

orange perithecia on bark

Arthonia

These inconspicuous crusts are common on bark,with at least 29 species reported from California,differing mainly in ascocarp appearance and sporetype (Grube 2008 [2007]) The ascocarps may becircular dark spots ~ 1 mm in diameter, or lirellae,branched or dendroid, pruinose or not, depending onthe species The nearly globose asci are usuallywithin one or two layers of the surface of the bark,and contain eight spores With Grube’s key available,

it is worth collecting likely species of Arthonia Probably the most common Arthonia is A pruinata, a white pruinose crust common on live oak, Quercus

agrifolia Other species that are common in the Santa

Barbara area are A lecanactidea, A pinastri, A.

sanguinea, and A tetramera In north coastal

counties, Arthonia ilicina occurs on alder

Several species of Arthonia were found by

Williams & Sillett (2007) in Humboldt County, but

the species reported (Arthonia arthonioides, A.

BULLETIN OF THE CALIFORNIA LICHEN SOCIETY 16 (1), 2009 Tucker – New Reports

leucopellaea, A stellaris), most of which would be

new to CA, are probably incorrect The specimens

respectively key to more likely species reported by

Grube (2008 [2007]) in the recent Sonoran Lichen

Flora as Arthonia pinastri, A glaucella, and A.

pyrrhuliza I’ve found A pinastri on pine, Quercus

agrifolia, Populus trichocarpa, and sycamore in

Santa Barbara County

Grube (2008 [2007]) has sunk the genus

Arthothelium (having muriform spores) in Arthonia

(mostly septate spores) He does not consider spore

type sufficient to separate these two genera A

common California Arthonia with muriform spores is

Arthonia beccariana The ascocarps are circular

black superficial spots on twigs The spores are 18-23

µm long I have found Arthonia beccariana on

cultivated Aralia and Beaucarnea (ponytail palm) in

Santa Barbara County and on Populus sp on the

campus of the University of California, Santa Cruz.

Another related species, Arthothelium norvegicum

Coppins & Tønsberg, was discovered on Vaccinium

ovatum twigs (Tønsberg & Williams 2006, citing C.

Williams 313, Humboldt Co collection; Williams &

Sillett 2007) It has not been transferred officially to

Arthonia so remains in Arthothelium for the time

being The spores are much larger than those of

Arthonia beccariana.

Candelaria pacifica Westberg

Reference to recent issues of CALS Bulletins,

summarized by Carlberg & Doell (2009), indicates

that most collectors are not yet recognizing that our

most common Candelaria on bark may be C.

pacifica Candelaria concolor has been widely

accepted as the common species across the U S., but

Westberg & Nash (2002) recognized that C pacifica

is very common on the west coast Most of my

collections, from Amador and Solano County in

central CA to those in Santa Barbara County, key to

C pacifica in Westberg’s key (2002) C pacifica is

characterized by having soredia below the tips, no

lower cortex, and by having 8-spored asci

Candelariella antennaria Räs., C biatorina M.

Westb., & C lutella (Vain.) Räs

Species of Candelariella on bark have bright

gold-colored crustose thalli, often with apothecia but

are generally small and overlooked Westberg’s key

(2004) reveals that there are several species on bark

that should be looked for in California Among these,

Candelariella biatorina (Westberg 2007a; Westberg

& Nash 2008 [2007]) is easily identified by its

biatorine apothecia (lacking a margin), as shown in a

color photograph in Sonoran Flora, Vol 3 It is found

on conifer bark or wood Two other species withlecanorine apothecia (having a margin) include

Candelariella antennaria with yellow apothecia on a

gray thallus and 8-spored asci, and C lutella with

yellow apothecia and thalli, and 24-32 spores perascus Three articles by Westberg (2007a, 2007b,

2007c) comprise a monograph of Candelariella for

the western U S and Mexico

Coenogonium luteum (Dicks.) Kalb & Lücking.

This species was formerly called Dimerella

lutea, and is easily recognized by the usually vivid

green crust on bark and pale orange apothecia, less

bright than a Caloplaca The only recent published

primary report for CA is by Judy Robertson (2002a)from Monterey County; I’ve found it in the pygmyforest in Monterey County, on Monterey cypress onthe Monterey peninsula, and on madrone on thecampus of University of California, Santa Cruz, on a

CALS foray (Tucker et al 2004) A second species,

Coenogonium pineti (Ach.) Lücking & Lumbsch,

has been reported only once in CA, near Larkspur inMarin County by Albert Herre in 1943; I haveverified the specimen, now in the Field Museumherbarium in Chicago

Hypogymnia gracilis McCune

This species was recognized and published byBruce McCune (2002) He identified many

specimens labeled H imshaugii from the Santa Barbara Botanic Garden herbarium as H gracilis,

based on the white internal surface and small regularholes on the lower side It is worth checking

specimens of H imshaugii, the most common

Hypogymnia in California, for unrecognized

collections of H gracilis Some records of H.

gracilis include D Keil, s.n., Los Osos, San Luis

Obispo Co., Nov 1955, OBI; T Nash 29969, Hastings Natural History Reserve, Monterey Co., ASU; S.

Tucker 37576, trail to East Peak, Mt Tamalpais,

Marin Co., SBBG

Menegazzia subsimilis (H Magn.) R Sant

This species resembles the more common M.

terebrata of the north-coastal flora except that it has

stalked soralia Looking through specimens labeled

M terebrata in herbaria will probably turn up one or

two of M subsimilis that had not been recognized Williams & Sillett (2007) found M subsimilis, a new report for California, in the tops of Sequoia trees in

Humboldt County

Pseudocyphellaria perpetua (Miadlikowska et

BULLETIN OF THE CALIFORNIA LICHEN SOCIETY 16 (1), 2009 Tucker – New Reports

al 2002) was also found by Williams & Sillett

(2007) It is a foliose macrolichen, resembling P.

crocata except that P perpetua has a yellow medulla

(versus white in P crocata), and mostly marginal

soralia (laminal as well as marginal in P crocata)

Punctelia

Five species of the foliose lichen Punctelia can

be found in California, with P perreticulata being the

most common on bark (Egan & Aptroot 2004) Most

species have white dots or pores (pseudocyphellae)

on the upper surface Reports of P subrudecta in CA

are misidentifications of P perreticulata The

photograph in Brodo et al (2001, p 609), while

labeled as P subrudecta, is P perreticulata On rock

one finds Punctelia stictica, common on boulders on

Mt Tamalpais in Marin County, and P punctilla, on

the Channel Islands and mainland of Santa Barbara

County, at the Santa Barbara Botanic Garden

Punctelia borreri and P ulophylla also have been

reported in the state

L ITERATURE C ITED

Brodo, I.M., S.D Sharnoff, & S Sharnoff 2001

Lichens of North America Yale University

Press, New Haven

Carlberg, T & J Doell 2009 California lichens by

county compiled from field trip reports in the

Bulletin of the California Lichen Society

Bulletin of the California Lichen Society 15 (2):

30-42

Egan, R.S., & A Aptroot 2004 Punctelia, pp

431-436, in T H Nash III, B D Ryan, P Diederich,

C Gries, & F Bungartz (eds.), Lichen flora of

the Greater Sonoran Desert region, Vol 2

Lichens Unlimited, Arizona State University,

Tempe

Esslinger, T.L 2008 A cumulative checklist for the

lichen-forming, lichenicolous, and allied fungi of

the continental United States and Canada North

Dakota State University, http://www.ndsu

.nodak.edu/instruct/esslinge/chcklst/chcklst7.htm

(posted 18 January, 2008), Fargo, N.D

Esslinger, T.L, & R.S Egan 1995 A sixth checklist

of the lichen-forming, lichenicolous, and allied

fungi of the continental United States and

Canada The Bryologist 98: 467-549

Grube, M 2008 (2007) Arthonia, pp.39-61, in T H.

Nash III, C Gries, & F Bungartz (eds.), Lichen

flora of the Greater Sonoran Desert region, Vol

3 Lichens Unlimited, Arizona State University,

Tempe

McCune, B 1997 Ptychographa, a lichen genus

new to North America The Bryologist 100: 240

239-McCune, B 2002 Hypogymnia, pp 228-238, in T.

H Nash III, B D Ryan, C Gries, & F Bungartz(eds.), Lichen flora of the Greater SonoranDesert region, Vol 1 Lichens Unlimited,Arizona State University, Tempe

Miadlikowska, J., B McCune, & F Lutzoni 2002,

Pseudocyphellaria perpetua, a new lichen from

western North America The Bryologist 105: 10

1-Robertson, J 2002a Pygmy Forest field trip,Mendocino County, March 16, 2002, and list ofmacrolichens of the Pygmy Forest Bulletin ofthe California Lichen Society 9 (1): 8-12.Spribille, T., C.R Björk, S Ekman, J.A Elix, T.Goward, C Printzen, T Tønsberg, & T Wheeler

2009 Contributions to an epiphytic lichen flora

of northwest North America: 1 Eight newspecies from British Columbia inland rainforests The Bryologist 112(1): 109-137

Tucker, S.C., J Robertson, & S Altermann 2004.Lichen foray on the campus of University ofCalifornia, Santa Cruz, May 15-16, 2004.Bulletin of the California Lichen Society 11(2):48-53

Tucker, S.C & B.D Ryan 2006 Revised catalogue

of lichens, lichenicolous and allied fungi inCalifornia Constancea 84 (University ofCalifornia, Berkeley) Online (http://ucjeps.berkeley.edu/constancea/84/)

Westberg, M 2004 Candelariella, pp 46-53, in T H.

Nash III, B D Ryan, P Diederich, C Gries, & F.Bungartz (eds.), Lichen flora of the GreaterSonoran Desert region, Vol 2 LichensUnlimited, Arizona State University, Tempe

Westberg, M 2007a Candelariella (Candelariaceae)

in western United States and northern Mexico:the species with biatorine apothecia TheBryologist 110(3): 365-374

Westberg, M 2007b Candelariella (Candelariaceae)

in western United States and northern Mexico:the polysporous species The Bryologist 110 (3):375-390

Westberg, M 2007c Candelariella (Candelariaceae)

in western United States and northern Mexico:the eight-spored, lecanorine species TheBryologist 110(3): 391-419

Westberg, M., & T.H Nash 2002 Candelaria, pp 116-118, in T H Nash III, B D Ryan, C Gries,

& F Bungartz (eds.), Lichen flora of the Greater

BULLETIN OF THE CALIFORNIA LICHEN SOCIETY 16 (1), 2009 Tucker – New Reports

Sonoran Desert region, Vol 1 Lichens

Unlimited, Arizona State University, Tempe

Westberg, M., & T.H Nash 2008 (2007)

Candelariella, pp 378-380, in T H Nash III, C.

Gries, & F Bungartz (eds.), Lichen flora of the

Greater Sonoran Desert region, Vol 3 Lichens

Unlimited, Arizona State University, Tempe

Williams, C.B., & S Sillett 2007 Epiphyte

communities on redwood (Sequoia

semper-virens) in northwestern California The

Bryologist 110 (3): 420-452

Williams, C.B., & L Tibell 2008 Calicium

sequoiae, a new lichen species from

north-western California, U S A The Lichenologist40:185-194

BULLETIN OF THE CALIFORNIA LICHEN SOCIETY 16 (1), 2009 Perlmutter – Basic Lichenology

Basic Lichenology 2: Reproduction

Gary B PerlmutterUniversity of North Carolina HerbariumUniversity of North CarolinaChapel Hill, NC 27599-3280lushik aol.comLichens are incredibly diverse organisms, with a

documented catalogue of over 4400 species in North

America (Tucker & Ryan 2006) and more being

discovered all the time They are diverse in a variety

of characters, such as thallus type, substrate

preference and pollution tolerance One set of

characters that holds great variety involves modes of

reproduction Structure and anatomy of lichen

reproductive organs are also key toward lichen

identification

Lichens can reproduce vegetatively or sexually,

and often do both Vegetative means can involve

specialized organs such as soredia or isidia, or simply

through thallus fragmentation Sexual reproduction

typically involves the fungal partner only, and the

variety of fruiting bodies or ascomata (sing ascoma)

is mind-boggling Not only that, but their internal

anatomy such as tissue arrangement, spore sacs

(known as asci) and the spores themselves vary quite

a bit across species and higher orders of taxonomy

like genus, family, etc Overall, lichens of all types

can have any of these structures, and often in

combination Reproductive traits are usually species

specific, thus their usefulness in identification

Vegetative Reproduction

Soredia are minute dispersal packets consisting

of a few algal cells wrapped by fungal threads, and

often appear as pale granules Their placement on the

lichen thallus varies, and depending on species, they

can appear on the thallus surface, on lobe tips,

margins or in patches of broken cortex known as

soralia In some loop lichens (Hypotrachyna), the

thallus can form pustules that break open to release

soredia; these are known as schizidia Finally, the

soredia of some crustose lichens can even comprise

the entire thallus These latter lichens, such as those

in the genus Lepraria, are known as dust lichens

Isidia are small, finger-like extensions of the

thallus, which break off and disperse in the wind

They have a cortex, medulla and algal layer, and can

be globular, cylindrical or branched Many rock

shield lichens (Xanthoparmelia) use isidia to

propagate themselves Both soredia and isidia are

best seen with a hand lens, but soralia can be seenwith the naked eye

Fragmentation is often employed by lichens thatare brittle and tend to break apart when dry, like

many reindeer lichens (Cladonia spp formerly known as Cladina) One unusual microfoliose lichen

found in the tropics and eastern North America that

uses fragmentation exclusively is Flakea papillata.

Its minute lobes are dichotomously branched andvery thin, and thus flake off (hence the name).Lacking any reproductive organs, this species was notknown to be a lichen until the 1990s and went by thename “The Thing” as bryologists and lichenologistsstruggled to classify it (Perlmutter 2006) Only recent

molecular analysis has placed F papillata into a

lichen family (Muggia et al 2009)

Asci, Spores and the Fungal Sexual Life Cycle

While the vegetative propagules serve toreproduce the lichen as a whole organism, it does notleave the opportunity for genetic material to mix andproduce offspring that are genetically different Inlichens only the fungal partner reproduces sexually(the photobiont cells merely divide), and the wayfungi “do it” is distinct from that of plants andanimals, earning their placement in a separateKingdom In a word, fungal sex is weird

For starters, the thallus is haploid, meaning thatonly one set of chromosomes exist in any given cellnucleus We animals are diploid, with two sets ofchromosomes per nucleus, and only our eggs andsperm are haploid Plants are also diploid, but theyinvolve an alternation of generations with haploidstages of their life cycle growing as multicellularorganisms, largely hidden in cones, flowers or freelygrowing on the soil as in mosses and liverworts.From our animal viewpoint, fungi can be seen asbackwards in their lifecycle But it gets even stranger

The life cycle begins with a spore (haploid, or n),

which germinates into a hypha This hypha splits andgrows, captures an alga or cyanobacterium, and thecombination of partners triggers the fungus toproduce a lichen thallus When two adjacent thallimeet, they may merge Because of this merging, the

BULLETIN OF THE CALIFORNIA LICHEN SOCIETY 16 (1), 2009 Perlmutter – Basic Lichenology

concept of the individual is not really applicable to

lichens, for a single thallus can represent one

individual or an entire colony all merged into one

form That’s why we call “individual” lichens

“thalli” When a thallus matures, ascomata develop

and this is where the fungal sex happens

In fungi there is no male and female, but rather

type “a” and type “b” Another strange thing with

fungi is that their cells are multinucleated, resulting

from the breakdown of cell walls In the base of a

developing ascoma, a hypha from one of the types

develops into a multinucleated ascogonium, while

that of the other type into an antheridium, containing

an equal number of nuclei as the former These two

microscopic organs merge and share nuclei, which do

not themselves merge, but instead lie side by side in a

stage known as dikaryon (literally “two nuts”, or n +

n) The dikaryotic stage is a character that both sac

and mushroom fungi share and have probably

evolved from a common ancestor

From the dikaryotic ascogonium, ascogenous

hyphae develop, each composed of cells containing

nuclei from the two parents The nuclei then merge,

becoming fertilized like egg and sperm The resulting

diploid (2n) structure becomes the ascus, or sac, from

which the Ascomycota or sac fungi get their name

Inside the ascus meiosis occurs, mixing the genetic

material from the parent nuclei and producing four

daughter cells (n) These cells undergo one mitotic

division, resulting in the set of eight ascospores

typical of lichens Together the spores and the ascus

develop analogous to offspring inside a womb until

the spores are ejected from the ascus to be carried by

the wind

Anatomy of the Ascoma

Ascomata come in two major types: apothecia

(open cups) and perithecia (closed cups) Withinthese two types is a dazzling array of forms Tounderstand their variety we must describe theiranatomy

A typical ascoma, say that of a rim lichen

(Lecanora; Figure 1), has five tissues These are the

eiphymenium, hymenium, hypothecium, exciple andthalline rim The uppermost tissue of the cup is theepihymenium, which is made up of the tips of shortfungal threads known as paraphyses, and tips of asci.These tips are often swollen and in the case ofparaphyses sometimes branched Often there aregranules in this tissue that give the disk its color, andsometimes granules also lie on top, giving the disk afrosted look The outer granules are known as pruina;such a disk is described as pruinose

Under the eiphymenium lies the hymenium Animportant tissue, this is where the spores areproduced They develop in the asci, which aresupported in an upright position by the paraphyses.This is so that when spores are dispersed from theasci tips, they leave the ascoma More on asci andspore diversity below These organs take root in thehypothecium, a dense fungal tissue made up of highlybranched hyphae, including ascogenous hyphae.Surrounding these tissues is the exciple, whichfunctions as the outer wall When exposed, it usuallyappears as a thin rim the same color as the disk, such

as in many button lichens (Buellia) The exciple can

surround the disk like a cup, or just the base of it, orjust the sides and not extending below In peritheciathe exciple completely surrounds the ascoma, leavingonly a hole for spore release And surrounding theexciple is a thalline rim, composed of cortical,medullary and photobiont tissues The thalline rim, asthe name implies, is the same color as the thallus

Types of Ascomata

As noted above there are aplethora of ascomatal types,which vary in the compositionand arrangement of tissues justdescribed Many of these arefamily specific, and can bereadily observed in the field.Following is a tour of fruitingbody diversity

Arthonioid apothecia are

the simplest forms of apothecia(Figure 2A), with a poorlydeveloped exciple and nothalline rim Superficially theyare often very small, and canFigure 1

BULLETIN OF THE CALIFORNIA LICHEN SOCIETY 16 (1), 2009 Perlmutter – Basic Lichenology

vary from dot-like to star-like, but often are irregular

in shape These are typical of the Family

Arthoniaceae, also known as dot, comma or asterisk

lichens These lichens are crustose with very simple

thalli, and are considered a basal lichen branch

(Martin Grube, pers comm.)

Lecanorine apothecia (Figure 2B & 2C) are the

quintessential apothecia, with all tissues present and

fully developed They are the ascomata found on rim

lichens (f Lecanoraceae), shield lichens (f

Parmeliaceae) and others within the Order

Lecanoromycetes

Lirellae are elongated apothecia (Figure 2D),

often branched to starlike Lirellae are diagnostic of

the crustose script lichens (f Graphidaceae), also

known as graphids

Lecideine apothecia (Figure 2E) lack thalline

rims, but have prominent excipular rims The rims are

the same color as the disk, and are usually black (i.e.,

carbonized) In shape they are typically flat, but in

more mature apothecia of some species, the exciple is

overgrown by the hymenial tissues and appear

convex Button lichens (genus Buellia) and rock

tripes (Lasallia and Umbilicaria) are typical in

bearing these apothecia as well as many crustose taxa

that are often difficult to identify These latter crusts

bear the name “Little Black Dots”, not unlike the

“Damn Yellow Composites” of the sunflower family

(Asteraceae) that frustrate our botanist friends

Biatorine apothecia (Figure 2F) are similar to

lecideine ones, but are paler in color with the exciple

noncarbonized They are typical of crusts in the

genera Bacidia and Biatora.

Maezedia are a specialized type of ascoma with

tissues so reduced that only a mass of loose spores

are seen Maezedia are usually stalked and are

diagnostic of the minute “stubble lichens” (e.g.

Calicium, Chaenothecopsis), which are themselves

indicative of healthy environments

In thelotremoid or “double-walled” apothecia

the thalline wall is separated from the exciple and is

termed a columella (Figure 2G) These are

characteristic of the largely tropical family

Thelotremataceae, known as barnacle or volcano

lichens (Note: thelotremoid lichens have just been

recognized to lie within the Graphidaceae through

molecular analysis, but the taxonomic change has not

yet been made [Papong et al 2009].)

Pertusarioid apothecia – The thalline margin is

thick and wart-like, covering the apothecium with

only a pore (ostiole) for spore release (Figure 2H)

Warts can contain one to several apothecia, each

distinguished by its ostiole The crustose Wart

Lichens (f Pertusariaceae: Pertusaria) are

characterized by pertusarioid apothecia

Perithecia – These are a type of ascoma largely

buried in the thallus in which the exciple has nearlyfully enveloped the inner tissues, leaving only anostiole for spore release (Figures 2I & 2J) Whilefound in separate evolutionary lines of sac fungi, itappears that this ascoma type has evolved from theopen condition of apothecia (Liu & Hall 2004).Perithecia usually appear as convex black fruitingbodies with an ostiole at the apex, although there aredeviations in this body plan They may be separate orgrouped; the groupings may be in thalline tissue in astructure called the pseudostroma (as in

Trypethelium), or the perithecia may be fused to form

one exipular mass with several ostioles This lattercondition is known as compound perithecia and is

characteristic of Mycoporum Most perithecial

lichens are crustose and represent several families;

but some, such as stipplescales (Dermatocarpon), are

squamulose

Pycnidia and Conidia

Resembling small perithecia are pycnidia, shaped reproductive bodies that produce asexualspores called conidia Like ascomata, pycnidia serve

flask-to reproduce only the fungal partner of the lichen.Conidia are produced by the budding of specializedhyphae inside a pycnidium and can appear in largenumbers Appearing as tiny black dots, pycnidia can

be produced on lichens bearing apothecia orperithecia, and in the latter they are distinguished inthe field by their smaller size Internally, pycnidia aredistinguished by lacking ascomatal tissues, and theconidia are generally smaller than spores, one celleach and colorless, and far more numerous

Asci and Ascospores

Both the asci and the spores they contain varyfrom species to species, and can even distinguishgenera or goups of higher taxonomic order Asci canvary in size, shape and details of their tips, where thespores eject from In arthonioid genera such as

Arthonia and Arthothelium, these are balloon-shaped,

whereas those of other genera may be either shaped or cylindrical Asci may contain one orseveral layers of walls, which function in sporerelease The tips of asci are typically thickened, andthis thickened part is termed the tholus Parts of thetholus and the ascal walls variously stain blue withiodine, which is important in keying of many lichens,especially crusts As the ascus and the spores insidemature, pressure builds inside, until the tip breaks,

club-BULLETIN OF THE CALIFORNIA LICHEN SOCIETY 16 (1), 2009 Perlmutter – Basic Lichenology

Figure 2 Ascomata A) Arthonioid apothecia B) Lecanorine apothecia C) Lecanorine in cross-section D)Lirellae E) Lecidine apothecia F) Biatorine apothecia G) Thelotremoid apothecia H) Pertusarioid apothecia

I and J) Perithecia Photography by Gary B Perlmutter (A, B, D, E, G, and H) and Mikki McGee (C, F, I, and J).Figure printed in color on back cover